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Philippine Journal of Science 150 (S1): 281-297, Special Issue on Biodiversity ISSN 0031 - 7683 Date Received: 22 Sep 2020

Diversity of in Intertidal and Shallow-water Areas of Samal Island,

Maybelle A. Fortaleza*, Joemarie J. Lanutan, Junissa M. Consuegra, and Cleto L. Nañola Jr.

Coral Reef Resiliency and Ecology Studies Laboratory College of Science and Mathematics, University of the Philippines Mindanao Mintal, Davao City, Davao del Sur 8000 Philippines

Echinoderms are immediately observable around the coastal shores of Samal Island. Unfortunately, studies on diversity are lacking despite its relevance to coastal and marine ecosystems management. This study attempts to update the existing records of echinoderms in Samal Island, however, limited to the intertidal and shallow-water areas. Field surveys from January 2018 to January 2019 yielded a total of 30 echinoderm belonging to 17 families. Most of the echinoderms encountered belonged to Classes Asteroidea (sea stars, nine species) and Ophiuroidea (brittle stars, nine species), followed by Echinoidea (regular and irregular urchins, six species) and Holothuroidea (sea cucumbers, six species). Of the 1,015 individuals, sea urchins comprised the majority of the total individuals recorded (43.15%), succeeded by sea stars (30.44%), brittle stars (18.04%), and sea cucumbers (8.37%). Richest diversity was observed in Catagman with 23 echinoderm species recorded. This was followed by Camudmud (14 species), Pangubatan (eight species), Balet (seven species), Aundanao (six species), and Kaputian (three species). The five most commonly occurring species were the Echinometra mathaei (Blainville, 1825), typicus (Müller and Troschel, 1840), scolopendrina (Lamarck, 1816), Diadema setosum (Leske, 1778), and laevigata (Linnaeus, 1758). Differences in distribution and abundance may be attributed to the variability of habitats available across sites. An intensive biodiversity assessment is recommended to evaluate how the physicochemical parameters and extent of anthropogenic activities shape the diversity and distribution of echinoderms on the island. The current work also provides the first documentation of ophiuroid diversity and the first record of the brittle stars, Breviturma krohi (Stöhr, Boissin & Hoareau, 2013) and Ophiocoma cf. cynthiae (Benavides- Serrato and O’Hara, 2008) in Samal Island, Philippines.

Keywords: diversity, echinoderms, intertidal areas, Samal Island

INTRODUCTION unsurprising given that holothuroids and asteroids are highly regarded for their economic and aesthetic values. Echinoderms from Classes Holothuroidea and Asteroidea In contrast, studies related to Crinoidea, Echinoidea, are among the well-studied taxa in Mindanao, particularly and Ophiuroidea are relatively few. Crinoids are popular in Davao Gulf (Gamboa et al. 2004; Bos et al. 2008a) subjects in underwater photography; however, taxonomic and Sarangani Bay (Pitogo et al. 2018). This comes works concerning feather stars in the Philippines are *Corresponding Author: [email protected] limited (Clark 1909; Messing 2003; Arguelles et al.

281 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity

2010). Philippine studies on echinoids have focused eTrex20). Each site was characterized in terms of mostly on economically important sea urchins such as monsoonal exposure (Matsumoto et al. 2020), slope the gracious Tripneustes gratilla (Linnaeus, (Emery 1961), relief (Benkendorff and Thomas 2007), 1758) (Juinio-Meñez et al. 2008; Casilagan et al. 2013), and substrate type (Table 1). whereas ophiuroid studies date back to Domantay and Domantay (1966) and Domantay and Conlu’s (1968) Macro- surveys consisted of day and night works. Although echinoderms are among the most observations conducted once every full moon week conspicuous components of intertidal and shallow-water of the month. Two 50-m belt transects with two-meter areas, it is evident that reports on biodiversity or species observation width were laid on the intertidal (high, mid, composition are lacking (Bos et al. 2008a). and low) and shallow-water areas parallel to the shore. Sampling for the echinoderms included low tide reef Published works that provided diversity information walks, snorkeling in shallow-water areas, and inspection on some echinoderms in Davao Gulf include Bos et al. of the underside of rocks and crevices. All echinoderms (2008a) who documented 11 asteroid species from Samal encountered every 5 m were recorded and photographed. and Talikud Islands, Davao del Norte, and Sonico (2018) The corresponding habitat type where they were seen was who reported five regular sea urchin species from Punta also noted. Dumalag, Davao City. Other studies that were related to echinoderms of Davao Gulf have focused on the The list of echinoderms, common names, pooled relative population dynamics of the horned sea star abundance, and conservation status is reported in Table 2. nodosus (Linnaeus, 1758) (Bos et al. 2008b), population Following the criteria of Pearse (2009), the pooled relative biology of the sand-sifting sea star abundance of echinoderms was considered “scarce” (Müller and Troschel, 1840) (Bos et al. 2011), spatio- when only 1–4 individuals were found. “Common” was temporal distribution of the crown-of-thorns assigned to echinoderms with 5–25 individuals present (COTS) Acanthaster planci (Linnaeus, 1758) (Agustin and “abundant” when there were more than 25 individuals and Nañola 2013), and its management during outbreaks observed. The corresponding conservation status of (Bos et al. 2013), documentation of the gleaning, drying, echinoderms listed was based on the International Union and marketing practices of sea cucumbers (Subaldo 2011), for Conservation of Nature (IUCN) Red List of Threatened and the supply chains and cost-benefit analyses of its Species (IUCN 2020). fishery and trade in Davao del Sur (Shuck et al. 2013). All echinoderms were identified using published While there are a number of research efforts concerning literature (see Table 3). Initial species identifications echinoderms of Davao Gulf, most of these were only and those that were unidentified were verified by limited to particular echinoderm taxa. Our knowledge of experts through personal communication. Diagnoses of their overall diversity, including those coming from other the echinoderm species encountered in Samal Island, classes, remains insufficient. Moreover, it is important Philippines are provided in Table 3. Species descriptions to consider that previous works may have been done discuss the morphological characteristics that served as the but have remained unpublished, which unfortunately basis for their identification. Habitat refers to the actual adds to the existing research gap as far as documenting habitat where the echinoderms were observed, including the local marine biodiversity is concerned. The current additional information from published literature. Lastly, work provides a preliminary report on the diversity and the information on local distribution is based on published distribution of echinoderms limited to the intertidal and sources, although most echinoderms listed are commonly shallow-water areas of Samal Island. Documenting the encountered by tourists, recreational divers, or underwater local diversity is of utmost importance to strengthen the enthusiasts anywhere in the country. protection and management efforts in Davao Gulf as one of the marine key biodiversity areas in the country. RESULTS AND DISCUSSION A total of 30 echinoderm species belonging to 17 METHODS families were recorded from six intertidal and shallow- Six rocky intertidal areas located in Barangays Aundanao, water areas of Samal Island, Davao del Norte (Table 2). Balet, Camudmud, Catagman, Kaputian, and Pangubatan Most of the echinoderm species recorded came from the in the Island Garden City of Samal, Davao del Norte Classes Asteroidea (Figure 2) and Ophiuroidea (Figure 3) were visited monthly from January 2018 to January 2019 with nine species, followed by Echinoidea (Figure 4) and (Figure 1). The geographic coordinates of each site Holothuroidea (Figure 5) with six species. Of the 1,015 were recorded using a handheld GPS device (Garmin individuals observed, echinoderms from Class Echinoidea

282 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity

Figure 1. Map showing the six study sites in Samal Island, Davao del Norte, Philippines where the macro-invertebrate surveys were conducted. Map was created in R software (R Core Team 2018) using the shape file downloaded from the Philippine GIS Data Clearinghouse (retrieved on 10 Sep 2020 from http://philgis.org).

Table 1. Description of the six intertidal and shallow-water areas studied in Samal Island, Davao del Norte, Philippines. Habitat characteristics such as monsoonal exposure (Matsumoto et al. 2020), relief (Benkendorff and Thomas 2007), and substrate types are indicated. Study areas Coordinates Monsoonal exposure Site description Aundanao 7° 5' 24" N, 125° 46' 58.08" E Northeast Medium relief; slope is steep relative to other study areas; sandy, rubble. The upper is a creviced rock wall descending into boulders, rocks, and sandy-coral debris substrate in the lower intertidal and shallow-water area. Balet 7° 9' 54" N, 125° 45' 27" E Northeast High relief; gradual slope; the high intertidal zone has a boulder measuring 1–3 m high. This descends into a rock platform with crevices and tide pools. Camudmud 7° 9' 36" N, 125° 41' 25.44" E Southwest Low relief; gradual slope; the intertidal zone is mixed with sandy- and coral rubble substrates. Seagrass patches extend to seagrass beds in the low intertidal zone. Catagman 7° 3' 36'' N, 125° 40' 45.48" E Southwest Low relief; moderate slope; the high intertidal zone has a limestone boulder about 1–2 m in height. This descends into an extensive rocky reef platform with patches of seagrass and tide pools. Soft and sandy-coral rubble substrates are found from the low intertidal to shallow-water area. Kaputian 6° 57' 36" N, 125° 43' 32.52" E Southwest Medium relief; gradual slope; the high intertidal zone is a creviced boulder about 2–6 m height, gradually descending into a narrow rocky reef platform. The low intertidal zone is characterized by sandy and rocky substrates, with live corals in further shallow-water areas. Pangubatan 6° 55' 48" N, 125° 47' 2.76" E Northeast Medium relief; steep slope; the intertidal and shallow-water area of Pangubatan is similar to Aundanao, having a rock wall at the high intertidal zone that descends into a narrow sandy-rocky area.

283 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity

Table 2. List of echinoderms encountered in the six intertidal and shallow-water areas of Samal Island, Davao del Norte, Philippines. Pooled relative abundance is categorized into scarce (1–4 individuals), common (5–25 individuals), and abundant (more than 25 individuals) (Pearse 2009). Abbreviations used to represent the study sites are as follows: AUN – Aundanao, BAL – Balet, CAM – Camudmud, CAT – Catagman, KAP – Kaputian, PAN – Pangubatan. Class and family Scientific name Common name Pooled relative abundance IUCN conservation and occurrence status Class Asteroidea (Sea stars) 1. Acanthasteridae Acanthaster planci Crown-of-thorns starfish Common; CAT, CAM Not Evaluated (Linnaeus, 1758) 2. Archasteridae Archaster typicus Sand-sifting sea star Abundant; CAM, CAT, Not Evaluated (Müller and Troschel, 1840) PAN 3. Asteropseidae Asteropsis carinifera Sheriff-badge sea star Scarce; CAT Not Evaluated (Lamarck, 1816) 4. Blue sea star Abundant; BAL, CAM, Not Evaluated (Linnaeus, 1758) CAT, PAN 5. Ophidiasteridae Multi-pore sea star Scarce; CAT Not Evaluated (Lamarck, 1816) 6. Ophidiasteridae Nardoa tuberculata Mottled sea star Scarce; CAT Not Evaluated (Gray, 1840) 7. granulatus Granulated sea star Scarce; CAT Not Evaluated (Lütken, 1869) 8. Oreasteridae novaeguineae Cushion star Common; BAL, CAT, PAN Not Evaluated (Müller and Troschel, 1842) 9. Oreasteridae Chocolate chip sea star Common; CAM, PAN Not Evaluated (Linnaeus, 1758) Class Echinoidea (Regular and irregular urchins) 10. Cidaridae Phyllacanthus imperialis Imperial lance urchin; Scarce; CAT Not Evaluated (Tenison-Woods, 1878) Pencil urchin 11. Diadematidae Diadema setosum Long-spined sea urchin Abundant; AUN, CAT, PAN Not Evaluated (Leske, 1778) 12. Diadematidae Echinothrix calamaris Banded sea urchin Scarce; AUN Not Evaluated (Pallas, 1774) 13. Echinometridae Echinometra mathaei Rock-boring urchin Abundant; AUN, CAT, Not Evaluated (Blainville, 1825) KAP, PAN 14. latecarinatus Keeled heart urchin Abundant; CAM Not Evaluated (Leske, 1778) 15. Laganidae Laganum laganum Abundant; CAM Not Evaluated (Leske, 1778) Class Holothuroidea (Sea cucumbers) 16. marmorata Chalkfish Scarce; CAT Data Deficient (Jaeger, 1833) (Conand and Purcell 2013) 17. Holothuriidae fuscocinerea Ashen Scarce; CAM, CAT Least Concern (Jaeger, 1833) (Conand et al. 2013a) 18. Holothuriidae Holothuria leucospilota Black sea cucumber Abundant; AUN, BAL, Least Concern (Brandt, 1835) CAT, KAP, PAN (Conand et al. 2013b) 19. Holothuriidae Holothuria scabra Sandfish Scarce; CAT Endangered (Jaeger, 1833) (Hamel et al. 2013) 20. Synaptidae Synapta maculata Maculated synaptid Abundant; AUN, CAM, Not Evaluated (Chamisso and Eysenhardt, CAT, PAN 1821) 21. Synaptidae Synaptula sp. sea cucumber Scarce; CAM Not Evaluated (Örsted, 1849)

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Class and family Scientific name Common name Pooled relative abundance IUCN conservation and occurrence status Class Ophiuroidea (Brittle stars) 22. Hemieuryalidae Ophioplocus imbricatus No common name Common; BAL, CAT Not Evaluated (Müller and Troschel, 1842) 23. Breviturma krohi (Stöhr, No common name Scarce; CAM Not Evaluated Boissin and Hoareau, 2013) 24. Ophiocomidae Ophiocoma scolopendrina Lagoon Abundant; AUN, BAL, Not Evaluated (Lamarck, 1816) CAM, CAT, PAN 25. Ophiocomidae Ophiocoma cf. cynthiae No common name Scarce; CAM, CAT Not Evaluated (Benavides-Serrato and O’Hara, 2008) 26. Ophiocomidae Ophiomastix annulosa Chain-link brittle star Abundant; AUN, BAL, Not Evaluated (Lamarck, 1816) CAM, CAT 27. Ophiocomidae Ophiomastix janualis Black brittle star Scarce; CAT Not Evaluated (Lyman, 1871) 28. Ophiocomidae Ophiomastix pictum No common name Scarce; CAT Not Evaluated (Müller and Troschel, 1842) 29. Ophiolepis cincta cincta No common name Scarce; CAM Not Evaluated (Müller and Troschel, 1842) 30. Ophiomyxidae Ophiarachna incrassata Green brittle star Common; CAT Not Evaluated (Lamarck, 1816)

Table 3. Echinoderms from intertidal and shallow-water areas of Samal Island, Davao del Norte, Philippines. Species in asterisk are new records for Samal Island. Scientific name Species description, habitat preference, and distribution Acanthaster planci The crown-of-thorns starfish (COTS) has 8–21 arms (Moran, 1990) with long aboral spines commonly 15–30 mm in length (Clark and Rowe 1971). The color of A. planci individuals may vary from green, red, and grey to purplish-blue (Moran, 1990). The species was observed crawling on coral rubble and on mixed sandy-rocky substrates in Samal Island.

It is one of the widely distributed asteroids in the world (de Dios and Sotto 2015); however, COTS outbreak in the Philippines is sporadic. Sightings of A. planci were reported in Talikud Island (Bos et al. 2008a), Southern Leyte (de Dios et al. 2015), Misamis Oriental (Llacuna et al. 2016), Guimaras Island (Myoung et al. 2017), and Agusan del Norte (Walag et al. 2018). Archaster typicus The sand-sifting sea star has flattened arms (Clark and Rowe 1971) with short spines along its edges (de Celis 1980). They are light grey in color and usually covered in sand. They are found in sandy areas of Samal Island and are likely to be more abundant in areas with seagrass beds, as observed in Camudmud and Catagman. It is a microphagous feeder typically found in sand flats (Mukaiet al. 1986).

It is distributed along the Indo-Pacific region and observable between 0–60 m depths (Colin and Arneson 1995). Their occurrence was reported throughout the Philippines (Fisher 1919), particularly in Batangas (Pinto 1982), Marinduque (de Celis 1980), Talikud Island (Bos et al. 2008a), Samar (Galenzoga and Quinones 2014), Misamis Oriental (Llacuna et al. 2016), and Guimaras (Myoung et al. 2017). Asteropsis carinifera The body of A. carinifera is dark green with pale gray smears (Chao 1999). The body is smooth but it appears granular because of the presence of crystal bodies (Clark and Rowe 1971). It has a thick epidermis and conical spines that surround the margin (Clark and Rowe 1971). This species was observed in the low intertidal zone of Catagman during nighttime reef survey. They are known to be found in rocky areas (Chao 1999).

This species is distributed throughout the Indo-West Pacific region (Clark and Rowe 1971). Its occurrence inthe Philippines was only reported in Talikud Island (Bos et al. 2008a). Linckia laevigata The blue sea stars have five to six slender arms (Fisher 1919), which can be orange or blue in color depending on the depth where they are found (Williams and Benzie 1993). The arms are thick and slender with blunt tips (Clark and Rowe 1971). They have a thick epidermis and a heavily calcified skeleton (Yamaguchi 1975). This species was observed in all study sites except in Camudmud and Kaputian. It occupies diverse microhabitats (Yamaguchi 1977), from rocky to coral rubble at 0 m to 20 m depth (Magsino and Ravago-Gotanco 2002).

This sea star is also widely distributed in the Indo-West Pacific region (Yamaguchi 1977).L. laevigata is found throughout the Philippines (Fisher 1919), particularly in Kalayaan Islands (Magsino and Ravago-Gotanco 2002), Talikud Island (Bos et al. 2008a), Bohol (Libres 2015), Guimaras Island (Myoung et al. 2017), and Agusan del Sur (Walag et al. 2018).

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Scientific name Species description, habitat preference, and distribution Linckia multifora The multi-pore star has an irregular number and length of arms (Clark and Rowe 1971) as a result of autotomy (Rideout 1978). It has varying colors from red to grayish blue, depending on its location on the shore (Crawford and Crawford 2007). The L. multifora individual encountered during the survey was brownish to khaki in color. Similar to L. laevigata, L. multifora was also observed on rocky and coral rubble substrates submerged in water in Catagman.

This species is commonly found in coral reefs of tropical intertidal to subtidal areas (Rideout 1978; Crawford and Crawford 2007). The species is widely distributed in the tropical Indo-West Pacific region (Rideout 1978) and was previously reported by Bos et al. (2008a) in Talikud Island. Nardoa tuberculata The disc and arms of mottled sea stars are covered with short tubercles (Clark 1967), although their marginal plates are not obvious on the dorsal side (Clark and Rowe 1971). Their body color is highly variable, from brown-yellow to gray- brown or maroon to dark gray (Clark 1921). This species was observed on coral rubble substrate in the low intertidal zone of Catagman during nighttime reef walks. It is commonly found in reef flats (Marsh 1994).

This species is widely distributed in the Indo-West Pacific region (Fisher 1919). The occurrence of the species was reported in Talikud Island (Bos et al. 2008a), Misamis Oriental (Llacuna et al. 2016), and Guimaras Island (Myoung et al. 2017). Choriaster C. granulatus has a smooth body texture and developed arms that are short and stout (Clark and Rowe 1971). Although granulatus the body of C. granulatus appears smooth, it has a thick and calcified skeleton (Yamaguchi 1975). Yamaguchi (1975) added that they usually stay on fully exposed reef flats hence, the need for a more robust skeleton. Two distant individuals were observed on coral rubble in the low intertidal zone and shallow waters. This species commonly occurs in sand flats and among corals and (Bos et al. 2008b).

This species is widely distributed throughout the and Northwestern Pacific regions (Fisher 1919). Its occurrence in the Philippines was reported in Talikud Island (Bos et al. 2008a) and Agusan del Norte (Walag et al. 2018). Culcita The cushion star is an inflated asteroid with a pentagonal shape (Chao 1999). The marginal plates are thickened and the novaeguineae pore areas are irregular (Clark and Rowe 1971). Its color is highly variable, from dark purple to olive and yellow (Olliff 2013; Myoung et al. 2017). Five color morphs of C. novaeguineae were observed and recorded in Balet and Catagman. They were encountered up to the mid-intertidal zone, typically submerged in rocky tide pools. The species is reef- associated but it can also be found in sand flats (Purwati and Lane 2004).

It is found in the Indian Ocean and Western Pacific region (Purwati and Lane 2004). This species was reported locally in Talikud Island (Bos et al. 2008a), Guimaras Island (Myoung et al. 2017), and Agusan del Norte (Walag et al. 2018). Protoreaster The chocolate chip sea star has an elevated disc (Clark and Rowe 1971) and the body color varies from crimson, pink, nodosus light brown, beige, green, blue to white (Clark 1921; Chim and Tan 2012), although the individuals encountered in Samal Island were typically orange. They have dark-colored tubercles that extend from the central disc of the organism and along the median of each arm (Chim and Tan 2012). Solitary individuals were encountered in sandy-seagrass substrates of Camudmud, Catagman, and Pangubatan. This sea star is a common inhabitant of the shallow sandy and seagrass areas in the tropics (Scheibling and Metaxas 2008).

The chocolate chip sea star is widely distributed in the Indian and Pacific Oceans (Fisher 1919). The species was recorded in Talikud Island (Bos et al. 2008a), Bohol (Libres 2015), Misamis Oriental (Llacuna et al. 2016), Guimaras Island (Myoung et al. 2017), and Agusan del Norte (Walag et al. 2018). Phyllacanthus The test of P. imperialis is dark brown and the genital pore is not elevated (Clark and Rowe 1971). The primary spines imperialis are dark brown and granulated while the secondary spines are lighter in color (Filander and Griffiths 2017). They were typically observed hiding under the rocks or boulders in shallow-water areas of Catagman.

Global distribution of P. imperialis includes , Sri Lanka, Bay of Bengal, East Indies, North , Philippines, , South Japan, and South Pacific Islands (Clark 1954; Clark and Rowe 1971). This species was documented in Verde Island Passage (Mooi and Munguia 2014). Diadema setosum This sea urchin is identified by its long spines and overall dark coloration (Domantay and Conlu 1968). It also has five white spots on the interambulacral midline and an orange ring around the anus (Lessios and Pearse 1996). Larger D. setosum were observed aggregating in deeper waters and sandy bottoms of Aundanao while the smaller individuals were found along the low intertidal zone, hiding under the rocks or crevices.

This species is widespread from Indo-West Pacific, from Africa to (Mayr 1954). This species was previously reported in Verde Island Passage (Mooi and Munguia 2014), Misamis Oriental (Llacuna et al. 2016), Agusan del Norte (Walag et al. 2018), and Davao City (Sonico 2018). Echinothrix The spines of E. calamaris are variable in color but the specimen that we encountered had black and white banded spines, calamaris similar to Domantay and Conlu’s (1968) description. The test is greenish and the anal cone is large (Filander and Griffiths 2017). One individual was found in shallow waters of Aundanao, resting under the boulder. This species can be found in seagrass beds (Colin and Arneson 1995).

They are known from Polynesia to the Red Sea and East Africa (Colin and Arneson 1995). Occurrence of E. calamaris in the Philippines was reported in Verde Island Passage by Mooi and Munguia (2014) and in Davao City (Sonico 2018).

286 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity

Scientific name Species description, habitat preference, and distribution Echinometra The test of E. mathaei is globose, with thick, slender spines that have white ring base (Clark and Rowe 1971; Filander mathaei and Griffiths 2017). The tips of spines in someE. mathaei individuals are also white. Six color morphs (brown, red, pink, green, yellow, and white) of the E. mathaei species complex were observed in Balet and Catagman. Rock-boring urchins are observed hiding under the rocks and crevices during daytime. They are so-called ‘rock-boring urchins’ because of the grooved paths they create on rocky platforms (Colin and Arneson 1995).

General distribution of this species is in the Indo-West Pacific, from Africa to Hawaii (Mayr 1954). Occurrence of the rock-boring urchins was reported in Verde Island Passage (Mooi and Munguia 2014), Agusan del Norte (Walag et al. 2018), and Davao City (Sonico 2018). Brissus The test of B. latecarinatus is oval in shape and has sunken petals (Nigam and Raghunathan 2017). Live B. latercarinatus latecarinatus found in Camudmud are white to pale pink in color. Tests of B. latecarinatus are abundant in sandy areas of Camudmud, Samal Island. The test length ranges from 60–80 mm.

This species is distributed throughout the Indo-West Pacific (Mayr 1954). Presence of B. latecarinatus is reported in Verde Island Passage (Mooi and Munguia 2014). Laganum laganum Sand dollars from Family Laganidae are flattened and have well-developed petals (Clark and Rowe 1971; Mooi 1989). They are distinguished from Peronella based on five genital pores and longer periproct that is located between the mouth and posterior margin (Clark and Rowe 1971). Several tests of sand dollars are visible throughout the sandy areas of Camudmud. Live L. laganum burrow in the sand during daytime.

They are distributed from Bay of Bengal, East Indies, North Australia, Philippines, and South Pacific Islands (Clark and Rowe 1971). Bohadschia The body wall of B. marmorata is thick; the mouth is ventrally located (Domantay 1953) and the anus is large and almost marmorata dorsal (Purcell et al. 2012). The color of this sea cucumber species is variable (Reyes-Leonardo 1984) but the individual observed in Catagman has a cream-colored body with dark brown blotches. The species was encountered on a sandy substrate during a nighttime survey in the shallow-water areas of Catagman. They are nocturnally active and they prefer aerated sandy areas (Kerr et al. 2006). This species is widely distributed in the Indo-West Pacific region except Hawaii (Reyes-Leonardo 1984). Local distribution of this species includes Batangas (Reyes-Leonardo 1984), Zamboanga (Domantay 1953), Samar and Leyte (Gajelan-Samson et al. 2011), Davao del Sur (Subaldo 2011), Palawan (Jontila et al. 2014), Bohol (Libres 2015), Basilan (Jaafar et al. 2018), and Sarangani Bay (Pitogo et al. 2018). Holothuria Purcell et al. (2012) described H. fuscocinerea as the only sea cucumber with dark brown ring on its dorsally-located fuscocinerea anus. Their body is cylindrical and grayish-brown in color (Kerr et al. 2006; Purcell et al. 2012). This species was encountered on coral rubble substrates in Camudmud and Catagman. They are typically restricted to boulder cover, coral rubble and fragments (Reyes-Leonardo 1984), and forereef slope (Kerr et al. 2006).

This sea cucumber species has patchy distribution in the Indo-West Pacific region (Reyes-Leonardo 1984; Purcell et al. 2012). It is also reported to occur in Samar and Leyte (Gajelan-Samson et al. 2011), Palawan (Jontila et al. 2014; Dolorosa et al. 2017), and Sarangani Bay (Pitogo et al. 2018). Holothuria This holothuroid species can be distinguished from H. coluber with its darkly colored tentacles (Purcell et al. 2012). H. leucospilota leucospilota has a completely black and elongated body that extends from inside the crevices for support while feeding (Kerr et al. 2006). This species is typically observed in tide pools in low intertidal zone and in shallow-water areas.

It is one of the most widely distributed holothurian species found in the Indo-Pacific region (Purcell et al. 2012). This species was reported in Palawan (Jontila et al. 2014; Dolorosa et al. 2017), Basilan Province (Jaafar et al. 2018), Agusan del Norte (Walag et al. 2018), and Sarangani Bay (Pitogo et al. 2018). Holothuria scabra The body is subcylindrical in shape and has variable color in life but often white to grayish-green dorsally and whitish gray ventrally (Domantay 1953). They are wrinkled dorsally (Purcell et al. 2012) and flat on the ventral surface (Reyes- Leonardo 1984). This species was encountered on sandy-seagrass patches of Catagman. Gleaners in the area collect them for consumption. Their bodies are often covered in sand and are common inhabitants of shallow-water areas with sandy-seagrass substrates (Purcell et al. 2012).

This species is widely distributed in the tropical Indo-Pacific, except in Hawaii and further east to Fiji (Purcell et al. 2012). Locally, H. scabra was reported in Batangas (Reyes-Leonardo 1984), Zamboanga (Domantay 1953), Davao del Sur (Subaldo 2011), Samar and Leyte (Gajelan-Samson et al. 2011), Palawan (Jontila et al. 2014; Dolorosa et al. 2017), Basilan Province (Jaafar et al. 2018), Agusan del Norte (Walag et al. 2018), and Sarangani Bay (Pitogo et al. 2018).

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Scientific name Species description, habitat preference, and distribution Synapta maculata Synaptids are easily recognized with their snake-like appearance and dark bands that run their cylindrical body longitudinally (Domantay 1953). Color may vary from lighter to darker shades of brown alternated with black. Synaptids were found typically resting or crawling on sandy-seagrass substrates.

They occupy sandy habitats and they are widespread in the Indo-West Pacific area except in Hawaii (Reyes-Leonardo 1984). This species was also reported in Batangas (Reyes-Leonardo 1984), Zamboanga (Domantay 1953), and Palawan (Jontila et al. 2014). Synaptula sp. Several Synaptula species remain undescribed (Kerr et al. 2006). This synaptid was observed in a tide pool on a rocky substrate during nighttime survey in Camudmud. Morphologically, the species closely resemble S. reciprocans but records of this species outside the Red Sea are incorrect (G. Paulay, pers. comm.). Ophioplocus Disc diameter = 23.2 mm; O. imbricatus individuals have small radial shields and distinctive dark markings on the the imbricatus aboral side of the disc, which Boissin et al. (2016) described its appearance like one side of a soccer ball.

This species was encountered from the mid-intertidal zone to shallow-water areas of Balet and Catagman. They are usually hidden under the rocks and submerged in water. Clark and Rowe (1971) reported that this species is widespread in the Indo-West Pacific region. Breviturma krohi* Disc diameter = 24.1 mm; B. krohi has variegated to uniform dark brown color pattern and weakly ringed arm spines (Stöhr et al. 2013). They have coarse dorsal disc granulation (max. of 164 mm2) and a maximum disc diameter of 16 mm (Stöhr et al. 2013). The B. krohi observed was dark brown in color and mottled, with banded arm spines. All arms are dark brown dorsally and light brown ventrally. The oral shields appear a shade darker than the light brown ventral color of the arms. The species was observed in the shallow-water area of Camudmud, along the sandy and coral rubble substrates.

This species is widely distributed across the Indian and Pacific oceans (Stöhret al. 2013; Boissin et al. 2016), occurring on coral reefs between 0–30 m (Hoareau et al. 2013). Ophiocoma Disc diameter = 22 mm; Boissin et al. (2016) reported that they can be identified from other members of the “scolopendrina scolopendrina group” based on their color. They are dark brown and variegated dorsally but paler ventrally (Clark and Rowe 1971). They are typically found in tide pools hiding underneath the rocks and boulders. They sway their arms when submerged in water (Boissin et al. 2016).

Species from the Ophiocoma are reported to have pantropical distribution and are relatively abundant in the Indo- West Pacific region (Clark and Rowe 1971; Devaney 1978). Ophiocoma cf. Disc diameter = 8.9 mm (juvenile); This species can be distinguished from other members of the O. erinaceus complex cynthiae* by the absence of granulation orally and presence of two tentacle scales that surround the tentacle pore (Benavides- Serrato and O’Hara 2008; Boissin et al. 2016). This brittle star is black in color and the tube feet are light to dark grey, in contrast to bright red of O. erinaceus. The species were observed in the low intertidal zones of Camudmud and Catagman.

The species occurs in the Great Barrier Reef, Australia to Tonga (Benavides-Serrato and O’Hara 2008) and was reported in Mascarenes (Boissin et al. 2016) and Lakshadweep Archipelago (Parameswaran et al. 2020). Philippine records appear to be plotted somewhere in Mindoro and the Verde Island Passage (OBIS 2020). Ophiomastix Disc diameter = 21.9 mm; This species has two tentacle scales, several blunt spinules on its disc, and distinctive red-grey annulosa coloration (Clark and Rowe 1971; Devaney 1978). This brittle star hides itself under the rocks in shallow tide pools. They are usually spotted from the mid-intertidal to shallow-water areas of Aundanao, Balet, Camudmud, and Catagman.

Members of the genus Ophiomastix are found in the tropical Indo-West Pacific region (Devaney 1978). Devaney (1978) also examined the O. annulosa specimen collected by Domantay from Batangas. Ophiomastix Disc diameter = 17.1 mm; The disc is dark with several light-colored spinules and pointed arm spines (Devaney 1978). janualis The specimen encountered has uniform black color with yellow segments on its arms. This species was observed only in the shallow-water area of Catagman, particularly along coral rubble substrate. Clark (1921) reported it to be common in Porites, Pocillopora, and Acropora (Devaney 1978).

This species was reported to occur in the Northern Great Barrier Reef, Indonesia, Philippines, and Ryukyu Islands (Devaney 1978). Ophiomastix pictum Disc diameter = 23.0 mm; The disc is naked, having gray and tan lines that somehow form a unique pattern. Dorsally, there is a longitudinal line intersecting in each of the arms (Clark and Rowe 1971). Clark and Rowe (1971) described this as Ophiarthrum pictum until recently, O’Hara et al. (2018) transferred this species to genus Ophiomastix. This species was encountered from the low intertidal to shallow-water area hiding under the rocks.

Generally, Ophiomastix species are limited to the tropical Indo-West Pacific region (Devaney 1978).

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Scientific name Species description, habitat preference, and distribution Ophiolepis cincta Disc diameter = 9.3 mm; The disc is flat and has overlapping plates that appear to be symmetrically arranged (Boissin cincta et al. 2016). Dorsally, this brittle star species is greenish-gray in color and white on the oral side (Domantay and Conlu 1968). Fujita (2016) reported that it should be represented as Ophiolepis cincta cincta instead of Ophiolepis cincta, following the suggestion of Stöhr et al. (2020). This brittle star was observed in the low intertidal zone of Camudmud and Catagman.

This species is found throughout the Indo-West and Central Pacific regions (Clark and Rowe 1971; Fujita 2016). Fujita (2016) also observed O. cincta cincta in the intertidal area of Singapore Strait. Local record of O. cincta in the Philippines was reported by Domantay and Conlu (1968). Ophiarachna Disc diameter = 43.4 mm; This species has granulated radial shields and is uniformly green in color with grey spots that incrassata appear to form a pattern around the disc (Clark and Rowe 1971). This species was observed along the rocky reef platform in the low intertidal zone of Catagman.

This species occurs in Sri Lanka, Bay of Bengal, North Australia, Philippines, China, South Japan, and South Pacific islands (Clark and Rowe 1971).

Figure 2. Sea stars (Echinodermata: Asteroidea) from intertidal and shallow-water areas of Samal Island, Philippines: A) Acanthaster planci; B) Archaster typicus; C) Asteropsis carinifera; D) Linckia laevigata; E) Linckia multifora; F) Nardoa tuberculata; G) Choriaster granulatus; H) ; I) Protoreaster nodosus. Photos by MA Fortaleza (A–B, D–E, and G–H), JJ Lanutan (C and F), and FBIO Aquino (J).

(43.15%) were the most abundant, followed by Class The common names, pooled relative abundance, Asteroidea (30.44%), Ophiuroidea (18.04%), and lastly distribution, and conservation status of echinoderms are Holothuroidea (8.37%). Overall, Echinometra mathaei listed in Table 2. Of the six study sites, Catagman has the (Blainville, 1825) is the most abundant echinoderm across richest diversity with 23 species, representing 76% of all study areas, followed by Archaster typicus (Müller and the total recorded echinoderm species. This is followed Troschel, 1840), Ophiocoma scolopendrina (Lamarck, by Camudmud (14 species), Pangubatan (eight species), 1816), Diadema setosum (Leske, 1778), and Linckia Balet (seven species), and Aundanao (six species). laevigata (Linnaeus, 1758) (Figure 6). Kaputian was the least diverse with only three species

289 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity

Figure 3. Brittle stars (Echinodermata: Ophiuroidea) from intertidal and shallow-water areas of Samal Island, Philippines: A) Ophioplocus imbricatus; B) Breviturma krohi; C) Ophiocoma scolopendrina; D) Ophiocoma cf. cynthiae; E) Ophiomastix annulosa; F) Ophiomastix janualis; G) Ophiomastix pictum; H) Ophiolepis cincta cincta; I) Ophiarachna incrassata. Photos by MA Fortaleza (B, D, F–G, I) and JJ Lanutan (A, C, E, H).

Figure 4. Regular and irregular sea urchins (Echinodermata: Echinoidea) from intertidal and shallow-water areas of Samal Island, Philippines: A) Phyllacanthus imperialis, B) Diadema setosum, C) Echinometra mathaei, D) Echinothrix calamaris, E) Brissus latecarinatus, F) Laganum laganum. Photos by MA Fortaleza (A, C–F) and FBIO Aquino (B).

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Figure 5. Sea cucumbers and synaptids (Echinodermata: Holothuroidea) from intertidal and shallow-water areas of Samal Island, Philippines: A) Bohadschia marmorata, B) Holothuria fuscocinerea, C) Holothuria leucospilota, D) Holothuria scabra, E) Synapta maculata, F) Synaptula sp. Photos by MA Fortaleza (A, C, F), JJ Lanutan (B, D), and FBIO Aquino (E).

Figure 6. Pooled relative abundance of echinoderm species from six intertidal and shallow-water areas of Samal Island, Davao del Norte, Philippines.

291 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity recorded in the area. Most of the echinoderms listed were works of Benavides-Serrato and O’Hara (2008), Pineda- “Not Evaluated” by the IUCN Red List of Threatened Enríquez et al. (2014, 2017), Stöhr et al. (2012, 2013), Species (Table 2). Conservation status was only reported Boissin et al. (2016, 2017), and O’Hara et al. (2017, in four economically-important holothuroids – namely, 2018, 2019) have brought advancements in understanding Holothuria scabra (Jaeger, 1833) (Endangered; Hamel et ophiuroid phylogeny and diversity. al. 2013), H. fuscocinerea (Jaeger, 1833) (Least Concern; Conand et al. 2013a), H. leucospilota (Brandt, 1835) Recent Philippine studies on Ophiuroidea include the (Least Concern; Conand et al. 2013b), and Bohadschia occurrence of Macrophiothrix longipeda (Lamarck, 1816) marmorata (Jaeger, 1833) (Data Deficient; Conand and in Gusa, Cagayan de Oro (Walag and Canencia 2016) and Purcell 2013). investigation on prey items of Ophiothrix spp. (Müller and Troschel, 1840) in Nogas Island, Philippines (Segumalian The species description, habitat information, and global and et al. 2016). The current study has reported only nine local distribution of the 30 echinoderms listed are provided species coming from four families under the Order in Table 3. Two echinoderm species are considered as new . This accounted for a total of nine species, two records in Samal Island – namely, Breviturma krohi (Stöhr, of which are new records for Samal Island (see Table 2). It Boissin and Hoareau, 2013) and Ophiocoma cf. cynthiae is likely that the other brittle stars we encountered would (Benavides-Serrato and O’Hara, 2008). These newly augment our current list but the difficulty of identifying recorded species may have long existed in Samal Island or them just through photographs indicates insufficient probably in other rocky intertidal shores in the country but effort. Knowing that the Indo-Pacific region holds the have remained undocumented until the recent echinoderm most number of ophiuroid species (Stöhr et al. 2012), this survey in the area. only reflects the need for local biodiversity inventories with complementary collection efforts to update what we Global diversity records revealed that Asteroidea and previously understood from the works of Domantay and Ophiuroidea are among the most diverse classes of Domantay (1966), Domantay and Conlu (1968), and Clark echinoderms (Mah and Foltz 2011; Mah and Blake 2012). and Rowe (1971). The asteroids listed in Table 2 are members of the Order (Asteroidea: Superorder Valvatacea), which are Four of the six species listed in the Class Holothuroidea reported to be the most speciose and ecologically diverse are economically-important sea cucumbers. This is across all echinoderm taxa (Blake 1983; Mah and Blake represented by the high-valued sea cucumber Holothuria 2012). Clark and Rowe (1971) reported that 77% of scabra, which was reported to be exploited in the Indo- the species in the Indo-West Pacific are from the Order Pacific region for over 1000 years (Conand 2004). Valvatida (Blake 1983). The diversity in ecological roles Typically, the dried forms of sea cucumbers, including among valvatidans qualifies them as ideal study organisms Bohadschia marmorata are made into “trepang” or in many fields of research (Mah and Foltz 2011). The “beche-de-mer” (Domantay 1953). Moreover, the less feeding ecology of Asteropsis carinifera (Lamarck, preferred H. fuscocinerea and H. leucospilota are among 1816), Choriaster granulatus (Lütken, 1869), Culcita the many holothuroids studied in marine pharmacology novaeguineae (Müller and Troschel, 1842), Linckia (Pangestuti and Arifin 2017; Khotimchenko 2018). laevigata, and L. multifora (Lamarck, 1816) have been Although the latter sea cucumbers were assessed as Data widely studied (Yamaguchi 1975, 1977). Further, decades Deficient and Least Concern based on the IUCN Red List of research have been dedicated to COTS outbreak and criteria, respectively, understanding their biology and their impacts on reefs (Alcala 1976; Wakeford et al. 2008; regional population is vital for sustainable use (Conand Tanner 2017). Many sea stars have also been considered as 2004). sources of specific bioactive and steroid compounds such as the Archaster typicus (Yang et al. 2011), C. granulatus The sea cucumbers reported in this list were common (Ivanchina et al. 2018), and Protoreaster nodosus (Thao and readily verified by experts. However, it is of utmost et al. 2015). importance to examine their calcareous rings, ossicles, and spicules for a more robust identification (Clark and Ophiuroids from intertidal to hadal depths have Rowe 1971; Rowe and Doty 1977; Purcell et al. 2012). notable ecological roles and are interesting subjects in Similar consideration must be taken when attempting paleontological studies (Stöhr et al. 2012; Boissin et al. to identify Synaptidae, especially that they are hardly 2016). Stöhr (2012) reported that the ophiuroid diversity discernible based on morphology alone (Kerr et al. has increased continually over the years. Despite their 2006; Martins and Souto 2020). Many synaptids in the abundance and widespread distribution, estimation of their Philippines are typically reported as either Synapta global diversity poses a challenge because many are still maculata (Chamisso and Eysenhardt, 1821) or Euapta awaiting description and proper taxonomic identification godeffroyi (Semper, 1868), although color patterns are (Stöhr et al. 2012; Boissin et al. 2016). Fortunately, the already varying. Whether these are just color morphs or

292 Philippine Journal of Science Fortaleza et al.: Echinoderms of Samal Island Vol. 150 No. S1, Special Issue on Biodiversity probably unreported synaptid species demands proper Garden City of Samal, Davao del Norte for allowing us taxonomic identification to reveal actual diversity. to conduct the fieldwork. Similar to holothuroids, echinoids that are harvested for Special thanks are given to the following experts for food are central to local research efforts in the Philippines. assisting us in the identification and verification of the The regular and irregular urchins listed in the study were echinoderms listed in this manuscript: Arthur R. Bos not of economic importance. However, large Diadema (The American University in Cairo) for Asteroidea; Kier setosum catches are occasionally harvested for human Mitchel Pitogo (Department of Environment and Natural consumption (de Guzman et al. 2016). Some locals in Resources Provincial Environment and Natural Resources Aundanao were also observed collecting large Echinothrix Office South Cotabato), Remie Aurelio [University of calamaris (Pallas, 1774) for food. the Philippines (UP) Mindanao], and Gustav Paulay (University of Florida) for Holothuroidea; and Neil Edgar Echinoids are also ecologically diverse, which presents Dex P. Marza (UP Los Baños), Tania Pineda-Enríquez challenges in their phylogeny and classification (Kroh (University of Florida), Timothy O’Hara (Museum and Smith 2010). Regular echinoids are popular subjects Victoria), Sabine Stöhr (Swedish Museum of Natural among recreational divers and underwater photographers, History), and Emilie Boissin (Université de Perpignan) but much of these citizen science efforts lead to for Ophiuroidea. inaccurate identification (Mooi and Munguia 2014). On the other hand, irregular echinoids receive relatively Charles G. Messing (Nova Southeastern University) less attention despite their abundance and ecological provided relevant information and resources on Crinoidea. roles as bioturbators (Belaústegui et al. 2017). Mooi and We also thank Fritzie Bianca Isabelle O. Aquino, Geleena Munguia (2014) reported a total of 210 echinoid species Ysabel B. Ang, Jerby Anne M. Basilio, Merlene E. in the Philippines based on combined literature, museum Elumba, Jodi Eugenia Lourdes F. del Fierro, and Kevin collections, and resulting ground-truthing from their 2011 L. Labrador for the field assistance. Lastly, we thank the expedition. Similar to ophiuroids, it is likely that the reviewers for their valuable comments that improved our diversity of irregular urchins may be underestimated in manuscript. the area because they are relatively less popular subjects for research. Overall, these findings will be augmented if research on the diversity and ecology of echinoderms in general will be supported. 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